Fluctuating asymmetry as a biomarker of habitat fragmentation in an area-sensitive passerine,the Eurasian treecreeper (Certhia familiaris)

The research investigating fluctuating asymmetry (FA), a phenotypic proxy of developmental instability, as a potential early-warning biomarker of anthropogenic stress like habitat fragmentation has produced controversial results. We examined the influence of habitat fragmentation at the landscape-scale, divided into habitat amount and configuration, on feather length FA in the Eurasian treecreeper (Certhia familiaris), an area-sensitive old-growth forest passerine breeding in boreal forests that are currently under intense management. Our study included one tail and wing feather, measured in both sexes of the 14-day-old nestlings and adults. Habitat amount was measured as the proportions of suitable forest habitats and open unsuitable areas, mean patch isolation and nesting patch size whereas habitat configuration was measured as mean patch density, size and shape and edge density. We found only weak sex- and age-related associations between feather length FA and habitat fragmentation that explained just 4.9% of variance in FA. Habitat loss was associated with higher FA in males only while the habitats with low degree of configuration tended to be related to lower FA in adults only. From the biomarker perspective, FA may not thus hold a great promise as a strong and general indicator of habitat fragmentation.     

Fluctuating feather asymmetry in relation to corticosterone levels is sex-dependent in Eurasian treecreeper (Certhia familiaris) nestlings

Fluctuating asymmetry (FA) has been widely used as a stress-related phenotypic marker of developmental instability. However, previous studies relating FA to various stressful conditions have produced inconsistent results and we still lack quantitative individual-level evidence that high FA is related to stress in wild vertebrate species. We studied how baseline plasma levels of corticosterone predicted FA of wing and tail feathers in free-living Eurasian treecreeper (Certhia familiaris) nestlings. We found a sex-specific association between corticosterone levels and FA: high corticosterone levels were related to an increased FA in male but not in female nestlings. These results suggest that in treecreepers, FA may correlate with individual stress hormone levels, male developmental trajectory being potentially more sensitive to stress than that of the female.     

Effects of forest patch size on physiological stress and immunocompetence in an area-sensitive passerine, the Eurasian treecreeper (Certhia familiaris): an experiment

We manipulated the primary brood size of Eurasian treecreepers (Certhia familiaris) breeding in different sized forest patches (0.5-12.8 ha) in moderately fragmented landscapes. We examined the effects of brood size manipulation (reduced, control, enlarged) and forest patch size on physiological stress (heterophil-lymphocyte ratios; H/L), body condition and cell-mediated immunocompetence (phytohaemagglutinin test). Nestlings' H/L ratios were negatively related to forest patch area in control and enlarged broods, whereas no effects were found in reduced broods. The effects of forest patch area were strongest in enlarged broods, which had, in general, twofold higher H/L ratios than control and reduced broods. The elevated H/L ratios were positively related to nestling mortality and negatively correlated with body-condition indices suggesting that the origin of stress in nestlings was mainly nutritional. Cell-mediated immunity of nestlings was not related to brood manipulation or to forest patch size. Also, the H/L ratios of adults were not related to brood manipulation or forest patch size. In addition, parental H/L ratios and body condition were not related to nestling H/L ratios. Our results suggest that during the breeding period the deleterious effects of habitat loss are seen explicitly in growing young.     

Within territory abundance of red wood ants Formica rufa is associated with the body condition of nestlings in the Eurasian treecreeper Certhia familiaris

Studies on individual reproductive success in relation to interspecific competition between distantly related taxa are scarce. We studied whether the abundance of red wood ants Formica rufa -group is related to the breeding habitat selection, fecundity and offspring quality in the Eurasian treecreeper Certhia familiaris, an old-growth forest passerine. The nest-box occupancy data were gathered over a five-year study period, whereas the breeding performance analyses were based on a two-year data set. The abundance of wood ants, measured within 50  m around the nest-boxes, was not related to nest-box occupancy rate, fecundity or the physiological stress of nestlings. In contrast, the abundance of wood ants was negatively related to the offspring quality and life-history traits, such as lowered body mass, subcutaneous fat reserves, and tarsus length at fledging. Our results suggest that exploitative competition between distantly related taxa may have considerable and adverse influences on nestling quality as measured by body mass, subcutaneous, fat and tarsus length. Red wood ants may decrease the fitness of treecreepers as the lower body condition of nestlings has the potential to impair recruitment into the breeding population and, additionally, impair the future reproductive effort.     

Genetic variation among Corsican and continental populations of the Eurasian treecreeper (Aves: Certhia familiaris) reveals the existence of a palaeoendemic mitochondrial lineage

In this study we investigated the phylogenetics of the Eurasian treecreeper (Certhia familiaris), a forest passerine with a wide Palaearctic range including Corsica, using three mitochondrial genes and three nuclear introns, and its phylogeographic history using the COI gene. Our phylogenetic results, including eight of the ten sub‐species currently recognized, support the monophyly of C. familiaris with respect to its Indo‐Asian sister species C. hodgsoni. C. familiaris comprises two lineages that diverged during the mid‐Pleistocene (c. 1 Myr): one palaeoendemic lineage has an allopatric range nowadays restricted to the Corsica island and the Caucasus region whereas the second one, more recent and widespread, is distributed over most of Eurasia and in northern China. The most likely scenario that may explain such a pattern is a double colonization of the western Palaearctic from the eastern range of the species. During the middle Pleistocene period, a first lineage expanded its range up into Europe but did not persist through glacial cycles except in Corsica and the Caucasus region. Later, during the upper Pleistocene, a second lineage began to diversify around 0.09 Myr, spreading towards the western Palaearctic from a unique refuge likely located in the eastern Palaearctic [correction added on 6 March 2015 after first online publication: 0.9 Myr amended to 0.09 Myr]. Apart from C. f. corsa, our results do not suggest any distinct evolutionary history for other sub‐species previously described on morphological grounds in Europe. Our study highlights the important conservation value of the Corsican treecreeper and emphasizes the major role of mature pine forests in the evolution of endemic bird taxa in Corsica. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 134–153.     

Forest fragmentation is associated with primary brood sex ratio in the treecreeper (Certhia familiaris)

We studied the primary brood sex ratio of an old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), along a gradient of forest fragmentation. We found evidence that male nestlings were more costly to produce, since they suffered twofold higher nestling mortality and were larger in body size than females. Furthermore, the proportion of males in the brood was positively associated with the provisioning rate and the amount of food delivered to the nestlings. During the first broods, a high edge density and a high proportion of pine forests around the nests were related to a decreased production of males. The densities of spiders, the main food of the treecreeper, were 38% higher on spruce trunks than on pine trunks. This suggests that pine-dominated territories with female-biased broods may have contained less food during the first broods. The observation was further supported by the fact that the feeding frequencies were lower in territories with high proportions of pines. In the second broods, territories with a high forest patch density produced female-biased broods, whereas high-quality territories with a large amount of deciduous trees and mixed forests produced male-biased broods. Our results suggest that habitat quality as measured by habitat characteristics is associated with sex allocation in free-living birds.     

Prey depletion and foraging strategy in the Oystercatcher Haematopus ostralegus

Summary The responses of a population of Oystercatchers to their own depletion of their prey, the edible cockle Cerastoderma edule, have been examined in Strangford Lough, Northern Ireland., Cockle stocks were severely depleted each winter as a result of predation by Oystercatchers and about half the birds present in October in the main study area had left by March: the reduction was greater when the initial population of Oystercatchers was high than when it was low. Oystercatchers were initially widespread around the Lough but tended to aggregate by January into a few good sites. Within each site annual differences in the location of feeding effort were correlated with year to year variation in the location of second winter cockles.Within the feeding areas in a bay the birds apparently hunted in the short-term on the basis of expectation, ceasing to crop, the densest cockle beds once an average yield for the area has been removed; this yield was apparently estimated from the local density of the anvils on which captured cockles were broken open. This effect led to marginal areas being depleted of cockles more rapidly than high density areas, so that the birds gradually concentrated their hunting onto an increasingly restricted area of the bay. The birds initially took only second winter and larger cockles but by late winter cockle densities had fallen so much that smaller cockles were accepted: areas previously abandoned or unused for hunting were then economically viable because of the reduced expectations of the birds and were incorporated into the feeding areas.These results are seen as consistent with Royama's profitability theory.     

Notes on Individual and Sexual Bill Variation in the British Tree-creeper (Certhia familiaris Britannica Ridgway)

T he purpose of this short treatise on the bills of Certhia familiaris Britannica Ridgway is to point out an as yet unrecognized sexual character in the species.     

Constant Proportions of Mixed Singers in Tree Creeper Populations (Certhia familiaris)

Of 205 tree creeper males in 7 syntopic populations 69 (34%) were mixed singers. Of these, 17 sang complete short-toed tree creeper song, six sang incomplete and 46 sang only one element thereof. In three populations (Neuenburg, Wendhausen, Kenzingen) there was a dominance of mixed singers, in another population (Lorettowald) a large proportion was present. Only one mixed singer was found in each of the last three populations (Freiburg, Bodanrück, Seerücken). Three populations had high proportions of mixed singers between 1960 and 1970 and later in 1983 and/or 1984. Two other populations had only one mixed singer in both periods. A small isolated population of 4 to 5 tree creeper pairs (Lorettowald) with many mixed singers was surrounded by populations with low proportions of mixed singers (Bodanrück, Seerücken). The proportions of different forms of mixed songs in Neuenburg in 1983 and 1984 were almost identical to those in the 1960′s whereas highly significant changes occurred in Kenzingen. The pronounced differences in the songs of tree creepers and short-toed tree creepers probably do not result from contrast reinforcement. Mixed song is also probably not caused by character (vocal) convergence according to Cody (1969) and Dobkin (1979). Instead, it seems more probable that mixed song results from errors in copying, which are passed on to the next offspring by cultural transmission.     

Prey selection by molluscivorous cichlids foraging on a schistosomiasis vector snail,Biomphalaria glabrata

Summary This paper considers prey size selection by four molluscivorous cichlids feeding on the intermediate host snail of Schistosoma parasites, Biomphalaria glabrata. Haplochromis ishmaeli obtains its prey by crushing the snails between the pharyngeal jaws, whereas H. xenognathus, H. sauvagei and Macropleurodus bicolor apply both pharyngeal crushing and oral shelling. The fishes crushed significantly more snails with the highest reward in biomass per second of crushing. Oral shelling occurred far less often than pharyngeal crushing. Encounter rates with prey showed significant variations between different size classes of prey. The fish have no overall knowledge of snail availability in a tank. The probability that a snail will be eaten at encounter, calculated from the number encountered and the number eaten, reflects the prey size preference of the fish. Those snails with the highest biomass/crushing-time ratio had the highest probability of being crushed; observed and predicted prey size preferences corresponded well. Although for oral shelling the potential reward in biomass per second is of the same magnitude as for crushing, the probability of successful shelling is very low. Apparently the fish prefer prey with lowest risks.     

Effects of aphids on foliar foraging by Argentine ants and the resulting effects on other arthropods

Abstract.  1. Although interactions between ants and honeydew-producing insects have received considerable study, relatively little is known about how these interactions alter the behaviour of ants in ways that affect other arthropods. In this study, field and greenhouse experiments were performed that examined how the presence of aphids ( Aphis fabae solanella ) on Solanum nigrum influenced the foraging behaviour of Argentine ants ( Linepithema humile ) and, in turn, modified the extent to which ants deter larval lacewings ( Chrysoperla rufilabris ), which are known aphid predators.
2. A field experiment demonstrated that the level of foliar foraging by ants increased linearly with aphid abundance, whereas no relationship existed between the level of ground foraging by ants and aphid abundance.
3. In the greenhouse, as in the field, foliar foraging by ants greatly increased when aphids were present. Higher levels of foliar foraging led to a twofold increase in the likelihood that ants contacted aphid predators. As a result of these increased encounters with ants, lacewing larvae were twice as likely to be removed from plants with aphids compared with plants without aphids. Once contact was made, however, the behaviour of ants towards lacewing larvae appeared similar between the two experimental groups.
4. Argentine ants drive away or prey upon a diversity of arthropod predators and parasitoids, but they also exhibit aggression towards certain herbivores. Future work should attempt to quantify how the ecological effects that result from interactions between honeydew-producing insects and invasive ants, such as L. humile , differ from those that result from interactions between honeydew-producing insects and native ants.     

Selective foraging on woody plant species by the Eurasian beaver (Castor fiber) in Telemark, Norway

Beavers Castor spp. are generalist herbivores, feeding on the bark, shoots and leaves of woody plants, terrestrial herbs and forbs, ferns and aquatic vegetation. As central-place foragers, beavers move out from water to select and cut trees and vegetation, and then transport it back to their refuge. These terrestrial forays are energetically costly; therefore, beavers should concentrate their foraging activity near the central place and increase the degree of selectivity for specific sizes or species of food with increasing distance from the water. The aim of this study was to test the predictions of the central place and the optimal foraging theories for the food selection of the Eurasian beaver Castor fiber , and show the foraging preferences of the focal species in the boreal conifer forest zone of Europe. Foraging intensity by beavers and the abundance of woody species were surveyed in transects positioned randomly at seven beaver territories. In compliance with the central-place foraging theory, the foraging intensity declined with increasing distance from the river. Beavers fed preferentially on willows ( Salix ), rowan ( Sorbus ) and birches ( Betula ), although alders ( Alnus ) dominated their diet. Size selectivity showed similar patterns to previous North American studies, which were also carried out in habitats with predominantly small saplings. The probability of selection of small saplings dropped as distance increased, which is consistent with the predictions of optimal foraging models that larger prey items are more likely to be favoured with increasing provisioning distance.     

Commensal association of the common kingfisher with foraging Eurasian otters

Positive interactions between birds and mammals are a fascinating aspect of animal behaviour. Feeding associations may consist of local enhancement or facilitation, and in the latter case, of commensalism or mutualism depending on the benefits received by the facilitator. We report here on a previously undescribed feeding association between piscivorous birds and Eurasian otters Lutra lutra. In Spain, common kingfisher Alcedo atthis and grey heron Ardea cinerea were observed closely following foraging otters and benefited from feeding opportunities provided by these. Behavioural observations of otters in central Spain (28.4 hr; 19 days) revealed that an association with kingfishers occurred in 33% of otter foraging events (n = 92). Simultaneous observations in northern Spain (14.2 hr; 16 days) showed an association between otters and kingfishers or grey herons in 41.6% and 11.7% of otter foraging events (n = 77), respectively. The association probability between kingfishers and otters increased significantly when otters foraged closer to the shore and on small fish rather than other prey (crayfish or large fish). Birds fed on prey remain left by the feeding otters, on small fish captured by otters when these were satiated and playing, or on prey displaced by otters. Our observations are consistent with facilitation and commensalism: piscivorous birds gained feeding opportunities provided by the otters, with no apparent costs or benefits to the latter. Similar feeding associations have been described between other species of otters and piscivorous birds (kingfishers, herons, egrets, storks) in Asia, South America and Southern Africa, but had not yet been described in Europe. The occurrence of piscivorous bird–otter associations in different species and regions suggests that this commensalism may be often overlooked but widespread. We have shown that the association can be frequent and is context‐dependent, with benefits for associating birds depending on otters´ behaviour and targeted prey.     

Optimal foraging by the swallow (Hirundo rustica, L): Prey size selection

The foraging ecology of the swallow (Hirundo rustica) was investigated in the field and compared with predictions of optimal foraging theory. Prey items were selected primarily by size. The inclusion of an item in the diet depended on the absolute abundance of the high-ranking prey and not of the low-ranking ones. Small items of low profitability were, however, included when food was abundant and more small items were taken when they were relatively abundant. Energetics data are used to show that it is profitable in terms of net energy gain for the swallow to take a mixture of large and small items, whereas optimal foraging theory would predict exclusive specialization on large, high-ranking items. Other possible explanations for the inclusion of small items in the diet are discussed.     

Optimal foraging: food patch depletion by ruddy ducks

Summary I studied the foraging behavior of ruddy ducks (Oxyura jamaicensis) feeding on patchily distributed prey in a large (5-m long, 2-m wide, and up to 2-m deep) aquarium. The substrate consisted of a 4x4 array of wooden trays (1.0-m long, 0.5-m wide, and 0.1-m deep) which contained 6 cm of sand. Any tray could be removed from the aquarium and loaded with a known number of prey. One bird foraged in the aquarium at a time; thus, by removing a food tray after a trial ended and counting the remaining prey, I calculated the number of prey consumed by the bird. I designed several experiments to determine if ruddy ducks abandoned a food patch in a manner consistent with the predictions of a simple, deterministic, patch depletion model. This model is based on the premise that a predator should maximize its rate of net energy intake while foraging. To accomplish this, a predator should only remain in a food patch as long as its rate of energy intake from that patch exceeds the average rate of intake from the environment. In the majority of comparisons, the number of food items consumed by the ruddy ducks in these experiments was consistent with the predictions of the foraging model. When the birds did not forage as predicted by the model, they stayed in the patch longer and consumed more prey than predicted by the model. An examination of the relation between rate of net energy intake and time spent foraging in the food patch indicated that by staying in a patch longer than predicted, the ruddy ducks experienced only a small deviation from maximum rate of net energy intake. These results provided quantitative support for the prediction that ruddy ducks maximize their rate of net energy intake while foraging.     

Prey selection and foraging period of the predaceous rocky intertidal snail,Acanthina punctulata

Summary The diet and foraging period of the neogastropod Acanthina punctulata were investigated in order to test various aspects of recent optimal foraging strategy models. This intertidal snail is an actively searching predator which preys on snails and barnacles by boring a hole in the shell and rasping out the flesh. Unlike many gastropod predators, Acanthina drill its gastropod prey at a very specific location on the columella, the thickest portion of the shell. Acanthina's foraging period can be interpreted as a compromise between maximizing the energy obtained by feeding and minimizing risk of mortality from exposure to wave action. That foraging period minimizing risk of being dislodged by waves appears to be during low tide when the predators can be in shallow pools. However, prey cannot be captured and consumed during one low tide. Thus Acanthina must be exposed during some high tides, and its strategy appears to be to restrict movement while exposed. Thus search is not initiated during high tide, but drilling and prey consumption are continued during that time. A snail not drilling or consuming prey seeks the protection of crevices or large anemones during high tide. A model is presented to indicate the relative amounts of risk and net energy for Acanthina at successive low and high tides. Predictions from the model, e.g., minimizing search time to avoid being exposed for an additional high tide and no movement during high tide are supported by field data. Acanthina commences foraging at the beginning of low tide, searches initially for preferred prey, but if unsuccessful, settles for a less preferred prey and begins drilling this prey before the end of low tide. Drilling and ingestion of prey occur during the following high and sometimes low tides. These handling times take 95% of the total foraging time in the field, while search time takes only 5% (pursuit time is negligible). Drilling alone accounts for 48–70% of the total drilling and eating time. In the laboratory, drilling and eating time for littorine food ranged from 15–60 hrs per item. The time to drill and eat a littorine increases exponentially with prey length. Since handling and processing prey items represents such a large investment of time, Acanthina would be expected to be very selective with respect to choice of prey items. Electivity coefficients from field data suggest that littorines are preferred over barnacles. Acanthina in the laboratory optimizes the amount of biomass ingested per time by choosing larger littorines over smaller ones and by preferring the more readily drilled species.It is suggested that Acanthina obtains information about the range of prey available initially by encountering and evaluating quite a few prey before making a selection, but usually by comparing an item of prey encountered to the prey it recently ingested. This latter method should provide a basis for evaluating prey encountered and has the advantage of reducing search time, the total amount of time spent feeding and thus the high-tide time exposed to wave action.In a similar manner, the decrease in the level of acceptability of prey as search time increases represents a compromise between maximizing energy obtained and minimizing risk from mortality.     

Prey size-distributions and size-specific foraging success of Ambystoma larvae

Summary We examined how prey size-distributions influence size-specific foraing rate and food gain, i.e., food intake scaled to metabolic demands, in Jefferson's and small-mouth salamander larvae. Ambystoma jeffersonianum larvae sampled on 17 dates from a farm pond whose fauna was dominated by macrozooplankton and chironomid larvae were rarely gape-limited, and total volume of food in the stomach (VS) showed only a slight tendency to increase with larval size. Although 15 of 17 correlation coefficients of VS with larval size were positive, only 1 of 17 correlations were statistically significant, and body size explained only 8% of the overall variation in VS. Correlation coefficients of food gain and body size were positive in 9 cases and negative in 8, but only 3 were statistically significant.In contrast, Ambystoma texanum larvae in 42 samples taken from five sites dominated by macrozooplankton as well as relatively large isopods and amphipods were almost always gape-limited, and VS tended to increase markedly with larval size. 40 of 42 correlation coefficients of VS and larval size were positive, and 19 correlations were statistically significant. Body size in turn explained about 35% of the overall variation in VS. Correlation coefficients of food gain and larval size were positive in 32 of 42 samples, and 9 of 10 significant correlations were positive.When food is limiting and prey selection is not limited by gape, smaller larvae may grow as fast or in some cases faster than larger larvae because they are nearly as effective foragers, but have lower metabolic demands. Larger larvae may in turn grow faster than smaller larvae in environments which support a broad size spectrum of prey, particularly when gape limitations are highly disproportionate among size classes. The growth rate of larvae in one size class relative to another depends primarily on the extent to which increased foraging rate compensates for higher energy demands as body size increases. Size-specific foraging rate may in turn be strongly influenced by the prey size-distribution within a habitat. These relationships suggest that relative size is not always a good a priori predictor of exploitative competitive ability.     

Prey location and prey choice by the freshwater leech Erpobdella octoculata using foraging kairomones

1. The freshwater leech, Erpobdella octoculata , is a generalist predator feeding on prey organisms such as Tubifex spp., Chironomus spp. and Asellus aquaticus. Using different experimental designs, we studied the use of foraging kairomones by the leeches for prey location.
2. Leeches were attracted to living as well as to freshly killed larvae of Chironomus sp., to Tubifex sp., and to A. aquaticus offered in Petri dishes covered with gauze. The leeches also reacted to an extract of macerated Tubifex sp. presented in agar.
3. Using ion exchange chromatography, the presence of histidine and glutamic acid was demonstrated in water contaminated by living larvae of Chironomus sp. and Tubifex sp. Agar blocks containing a synthetic mixture of these compounds at concentrations above 5 mg mL⁻¹ were attractive to the leeches.
4. Choice tests showed that leeches preferred chemical signals from Tubifex sp. over larvae of Chironomus sp. No difference was found between chemicals from Tubifex sp. and A. aquaticus , and A. aquaticus and larvae of Chironomus sp.
5. The results demonstrate that E. octoculata uses specific foraging kairomones in searching for prey and indicate that amino acids serve as foraging kairomones.